Abstract
Solanaceae is one of the most diverse families in the Americas, particularly in Argentina where it represents the fourth family in terms of species number. Although checklists for most South American countries have been published, some are outdated and there has been no analysis of Solanaceae diversity at country level. We present an updated summary of Solanaceae diversity in South America, an analysis of its distribution in Argentina, and preliminary conservation assessments for all species endemic to Argentina. Regression analyses were used for evaluating the ratio between taxa/area and endemic/total species, multivariate ordering methods were used to analyze the relationships between Argentine ecoregions, and the IUCN criteria were applied for conservation assessments. Results show that Solanaceae comprises 1611 species in South America. The highest diversity is in Peru, which, together with Ecuador, possesses more diversity than expected for the area; Chile and Brazil have the greatest percentage of endemic species. In Argentina, the Chaco ecoregion hosts the highest number of taxa, but largest number of endemic species is found in the Monte ecoregion. According to the IUCN criteria, 28 endemic species from Argentina are considered threatened. We discuss South American countries and Argentine ecoregions in terms of conservation priorities.
Key words
Argentina; conservation status; distribution; ecoregion; endemic species; South America
INTRODUCTION
The Solanaceae family includes 98 genera and approximately 2800 species (Dupin et al. 2017DUPIN J, MATZKE NJ, SÄRKINEN T, KNAPP S, OLMSTEAD RG, BOHS L & SMITH SD. 2017. Bayesian estimation of the global biogeographical history of the Solanaceae. J Biogeogr 44(4): 887-899.). It is distributed in all continents except Antarctica, with a preference for warm to tropical zones, from sea level to heights up to 5000 m and it inhabits many heterogeneous environments (Barboza 2013BARBOZA GE (Coord). 2013. Solanaceae. In: Zuloaga FO, Belgrano M & Anton AM (Eds), Flora Argentina, 1st ed., San Isidro: IBODA-IMBIV, CONICET 13: 1-350.). It includes small annual herbs (e.g. Leptoglossis Benth.) up to trees taller than 15 m (e.g. Duckeodendron Kuhlm.) (Hunziker 2001HUNZIKER AT. 2001. The genera of Solanaceae, 1st ed., Ruggell: A.R.G. Gantner Verlag K.-G, p. 1-500.) and it is well-known for its diverse reproductive structures (Knapp 2010KNAPP S. 2010. On “various contrivances”: pollination, phylogeny and flower form in the Solanaceae. Philos Trans R Soc London B Biol Sci 365(1539): 449-460.) as well as its varied chemical compounds (Eich 2008EICH E. 2008. Solanaceae and Convolvulaceae: Secondary metabolites: Biosynthesis, chemotaxonomy, biological and economic significance (a handbook), 1st ed., Berlin Heidelberg: Springer-Verlag, p. 1-637.). The Solanaceae species are very valuable to man mainly as food (Solanum tuberosum L., potato; Solanum lycopersicum L., tomato; Solanum melongena L., aubergine; Capsicum annuum L., bird pepper), ornamentals (species of Petunia Juss., Schizanthus Ruiz & Pav., Salpiglossis Ruiz & Pav., Browallia L., Brugmansia Pers.) and for their medicinal, poisonous or psychotropic effects (Nicotiana tabacum L., tobacco; Atropa bella-donna L., belladonna; Mandragora officinarum L., mandrake; Hyoscyamus niger L., henbane; Datura stramonium L., thorn apple; Cestrum parqui L’Her., duraznillo negro). Likewise, many taxa are used in biological studies as model organisms for experimentation e.g.: Nicotiana spp., Solanum spp., Petunia spp., Datura spp. (Gebhardt 2016GEBHARDT C. 2016. The historical role of species from the Solanaceae plant family in genetic research. Theor Appl Genet 129(12): 2281-2294.).
Solanaceae is among the 12 most diverse families in the Americas, the genus Solanum L. being the sixth in the number of species (Ulloa Ulloa et al. 2017ULLOA ULLOA C ET AL. 2017. An integrated assessment of the vascular plant species of the Americas. Science 358(6370): 1614-1617.), while South America represents both the ancestral area and the actual centre of diversity of the family (Olmstead 2013OLMSTEAD RG. 2013. Phylogeny and biogeography in Solanaceae, Verbenaceae and Bignoniaceae: a comparison of continental and intercontinental diversification patterns. Bot J Linn Soc 171(1): 80-102., Dupin et al. 2017DUPIN J, MATZKE NJ, SÄRKINEN T, KNAPP S, OLMSTEAD RG, BOHS L & SMITH SD. 2017. Bayesian estimation of the global biogeographical history of the Solanaceae. J Biogeogr 44(4): 887-899.). Argentina presents a vast richness of species; in fact, Solanaceae is the fourth family by number of species after Asteraceae (1502 spp.), Poaceae (1170) and Fabaceae (760), and is represented by 319 species (Barboza 2013BARBOZA GE (Coord). 2013. Solanaceae. In: Zuloaga FO, Belgrano M & Anton AM (Eds), Flora Argentina, 1st ed., San Isidro: IBODA-IMBIV, CONICET 13: 1-350., Del Vitto & Petenatti 2015DEL VITTO LA & PETENATTI EM. 2015. Sobre la presencia de Solanum rostratum (Solanaceae) en Sudamérica: una neófita tóxica de gran potencial como maleza. Rev Fac Cienc Agrar Univ Nac Cuyo 47(2): 109-121.).
Currently, species lists of vascular plants in Argentina (Zuloaga et al. 1999ZULOAGA FO, MORRONE O & RODRÍGUEZ D. 1999. Análisis de la biodiversidad en plantas vasculares de la Argentina. Kurtziana 27(1): 17-167.) and the South American Southern Cone (Zuloaga & Belgrano 2015ZULOAGA FO & BELGRANO MJ. 2015. The Catalogue of Vascular Plants of the Southern Cone and the Flora of Argentina: their contribution to the World Flora. Rodriguésia 66(4): 989-1024.) already exist. Nevertheless, neither an in-depth analysis of Solanaceae diversity nor preliminary conservation assessments applying the IUCN criteria for all species endemic to Argentina, have yet to be carried out.
The aims of this paper are: (i) to present an updated summary of the diversity of Solanaceae in South America and Argentina, related to its distribution, (ii) to assess the conservation status of all endemic species from Argentina and to identify the priority areas for planning regional conservation management in the country.
MATERIALS AND METHODS
Solanaceae diversity in South America
All South American countries were included in the diversity analysis. The species lists were compiled from floras and catalogues of the different countries (D’Arcy et al. 1993D’ARCY WG, HUNZIKER AT, BOHS LA, KEEL S, KNAPP S, MIONE T, NEE MH, RICK CM & SPOONER DM. 1993. Solanaceae. In: Brako L & Zarucchi JL (Eds), Catalogue of the flowering plants and Gymnosperms of Peru, 1st ed., Monogr Syst Bot Missouri Bot Gard 45: 1098-1137., 2007D’ARCY WG, BENÍTEZ DE ROJAS C & NEE MH. 2007. Solanaceae. In: Funk V, Hollowell T, Berry P, Kelloff C & Alexander SN (Eds), Checklist of the Plants of the Guiana Shield (Venezuela: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French Guiana), 1st ed., Contr US Natl Herb 55: 526-531., Short et al. 1999SHORT MJ, KNAPP S, ANDERSON GJ, CASTILLO R, MIONE T, NEE MH, SAWYER NW & SPOONER DM. 1999. Solanaceae. In: Jørgensen PM & León-yánez S (Eds), Catalogue of the vascular plants of Ecuador, 1st ed., Monogr Syst Bot Missouri Bot Gard 75: 900-918., Ulloa Ulloa et al. 2004ULLOA ULLOA C, ZARUCCHI JL & LEÓN B. 2004. Diez años de adiciones a la flora del Perú: 1993-2003. Arnaldoa Ed Especial 1-242., Rodríguez et al. 2006RODRÍGUEZ EF, VÁSQUEZ R, ROJAS R, CALATAYUD G, LEÓN B & CAMPOS J. 2006. Nuevas adiciones de angiospermas a la flora del Perú. Rev Peru Biol 13(1): 129-138., Hokche et al. 2008HOKCHE O, BERRY PE & HUBER O. 2008. Nuevo catálogo de la flora vascular de Venezuela, 1st ed., Caracas: Fundación Instituto Botánico de Venezuela, p. 1-859., Zuloaga et al. 2008ZULOAGA FO, MORRONE O & BELGRANO M. 2008. Catálogo de las Plantas Vasculares del Conosur (Argentina, Sur de Brasil, Chile, Paraguay y Uruguay). Online edition: http://www.darwin.edu.ar/Proyectos/FloraArgentina/fa.htm. Accessed on May/2020.
http://www.darwin.edu.ar/Proyectos/Flora...
, Knapp et al. 2011KNAPP S, ULLOA ULLOA C & MONTÚFAR R. 2011. Solanaceae. In: León-Yánez S, Valencia R, Pitman N, Endara L, Ulloa Ulloa C & Navarrete H (Eds), Libro rojo de las plantas endémicas del Ecuador, 2nd ed., Quito: Publicaciones del Herbario QCA, Pontificia Universidad Católica del Ecuador, p. 782-792., Barboza 2013BARBOZA GE (Coord). 2013. Solanaceae. In: Zuloaga FO, Belgrano M & Anton AM (Eds), Flora Argentina, 1st ed., San Isidro: IBODA-IMBIV, CONICET 13: 1-350., Nee 2014NEE MH. 2014. Solanaceae. In: Jørgensen PM, Nee MH & Beck S (Eds), Catálogo de las Plantas Vasculares de Bolivia. Monogr Syst Bot Missouri Bot Gard 127: 1203-1228., Orozco et al. 2015OROZCO CI , VÉLEZ JM, SARMIENTO Y, CANAL D, OREJUELA A, BELTRÁN G, BARBOZA GE, ALBA A & BERNAL R. 2015. Solanaceae. In: Bernal R, Gradstein SR & Celis M (Eds), Catálogo de plantas y líquenes de Colombia, Bogotá: Instituto de Ciencias Naturales, Universidad Nacional de Colombia. Available at http://catalogoplantasdecolombia.unal.edu.co/es/. Accessed on May/2020.
http://catalogoplantasdecolombia.unal.ed...
, Rodriguez et al. 2018RODRIGUEZ R ET AL. 2018. Catálogo de las plantas vasculares de Chile. Gayana Bot 75(1): 1-430., Flora do Brasil 2019aFLORA DO BRASIL. 2019a. Solanaceae in Flora do Brasil 2020 em construção. Jardim Botânico do Rio de Janeiro. Disponível em http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB225. Acessado em Maio/2020.
http://floradobrasil.jbrj.gov.br/reflora...
) and monographs of the genera when available. All taxonomic novelties up to April 2019 were included based on a web search with specific keywords (Solanaceae “new combination”, “new record”, “new species”, “sp. nov.” and “updated list”, https://scholar.google.com). Doubtful data were cross-checked against the databases Solanaceae Source (2019)SOLANACEAE SOURCE. 2019. A global taxonomic resource for the nightshade family. Available at http://www.solanaceaesource.org/. Accessed on May/2020.
http://www.solanaceaesource.org/...
and POWO (2019)POWO - PLANTS OF THE WORLD ONLINE. 2019. Facilitated by the Royal Botanic Gardens, Kew. Available at http://www.plantsoftheworldonline.org/. Accessed on May/2020.
http://www.plantsoftheworldonline.org/...
, and in all cases the final decision for species inclusion was based on specimens properly identified by Solanaceae specialists.
Linear regression analysis was used to determine the taxa-area ratio and the endemic species-species richness ratio for each country and the Guianas (French Guiana, Guyana and Suriname). When necessary, data were log-transformed to meet the assumptions of regression analysis. InfoStat v. 2016 (Di Rienzo et al. 2016DI RIENZO JA, CASANOVES F, BALZARINI MG, GONZALEZ L, TABLADA M & ROBLEDO CW. 2016. InfoStat version 2016. Grupo InfoStat, FCA, Universidad Nacional de Córdoba, Argentina. Available at http://www.infostat.com.ar/ . Accessed on May/2020.
http://www.infostat.com.ar/...
) software was used to perform all the statistical analysis.
Solanaceae diversity and distribution in Argentina
Diversity and distribution of Solanaceae in Argentina was analysed according to the ecological systems proposed by Josse et al. (2003)JOSSE C ET AL. 2003. Ecological systems of Latin America and the Caribbean: a working classification of terrestrial systems, 1st ed., Arlington: NatureServe, p. 1-47. and modified by Zuloaga & Belgrano (2015)ZULOAGA FO & BELGRANO MJ. 2015. The Catalogue of Vascular Plants of the Southern Cone and the Flora of Argentina: their contribution to the World Flora. Rodriguésia 66(4): 989-1024.. Eight ecoregions were considered: South Central Dry Andes, Atlantic Forest, Chaco, Monte, Wet Temperate Pacific, Pampas, Patagonia, and Yungas (Fig. 1).
Ecoregions of Argentina (modified from Zuloaga & Belgrano 2015ZULOAGA FO & BELGRANO MJ. 2015. The Catalogue of Vascular Plants of the Southern Cone and the Flora of Argentina: their contribution to the World Flora. Rodriguésia 66(4): 989-1024.). Photographs of typical Solanaceae species from each ecoregion are included.
Data from Documenta Florae Australis (2019)DOCUMENTA FLORAE AUSTRALIS. 2019. Available at http://www.darwin.edu.ar/iris/ .Accessed on May/2020.
http://www.darwin.edu.ar/iris/...
and GBIF (2019)GBIF - GLOBAL BIODIVERSITY INFORMATION FACILITY. 2019. Available at https://www.gbif.org/ . Accessed on May/2020.
https://www.gbif.org/...
databases were included in the analysis, complemented with information from specimen labels of the main herbaria of Argentina {BA, BAB, CORD, LIL, MERL, SI [Thiers continuously updatedTHIERS B. [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available at http://sweetgum.nybg.org/science/ih/. Accessed on May/2020.
http://sweetgum.nybg.org/science/ih/...
]} and monographs when available, i.e., Fabiana Ruiz & Pav. (Barboza & Hunziker 1993BARBOZA GE & HUNZIKER AT. 1993. Estudios en Solanaceae XXXIV. Revisión taxonómica de Fabiana. Kurtziana 22: 109-153.), Jaborosa Juss. (Barboza & Hunziker 1987BARBOZA GE & HUNZIKER AT. 1987. Estudios sobre Solanaceae XXV. Revisión de Jaborosa. Kurtziana 19(1): 77-153.), Lycium L. (Bernardello 1986BERNARDELLO G. 1986. Revisión taxonómica de las especies sudamericanas de Lycium (Solanaceae). Bol Acad Nac Ci Córdoba 57(3-4): 173-356.), Sclerophylax Miers (Di Fulvio 1961DI FULVIO TE. 1961. El género Sclerophylax (Solanaceae). Kurtziana 1: 9-103.), Solanum (Knapp 2013KNAPP S. 2013. A revision of the Dulcamaroid Clade of Solanum L. (Solanaceae). PhytoKeys 22: 1-432., Spooner et al. 2016SPOONER DM, ALVAREZ N, PERALTA IE & CLAUSEN AM. 2016. Taxonomy of wild potatoes and their relatives in Southern South America (Solanum sect. Petota and Etuberosum). Syst Bot Monogr 100: 1-240., Knapp et al. 2017KNAPP S, SAGONA E, CARBONELL AK & CHIARINI F. 2017. A revision of the Solanum elaeagnifolium clade (Elaeagnifolium clade; subgenus Leptostemonum, Solanaceae). PhytoKeys 84: 1-104.). When the identification of some specimens or their distributions were doubtful, Solanaceae experts were consulted.
The relationships between ecoregions were explored using principal coordinates analysis (PCoA) and a minimum spanning tree (MST). Jaccard’s similarity index (S) was calculated for the binary data of presence of taxa with the transformation (1-Sij)1/2.
Conservation status assessment of Argentine endemic species
The distribution for each endemic species was plotted using QGIS 2.8 (QGIS Development Team 2018QGIS DEVELOPMENT TEAM. 2018. QGIS Geographic Information System. Open Source Geospatial Foundation. Available at http://qgis.osgeo.org. Accessed on May/2020 .
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) and was based on georeferenced data of all the herbarium collections studied. Conservation status was assessed using the IUCN criteria version 3.1 (IUCN 2012IUCN - INTERNATIONAL UNION FOR CONSERVATION OF NATURE. 2012. IUCN Red List Categories and Criteria: Version 3.1, 2nd ed., Gland and Cambridge: IUCN, p. 1-32., IUCN Standards and Petitions Subcommittee 2017IUCN STANDARDS AND PETITIONS SUBCOMMITTEE. 2017. Guidelines for Using the IUCN Red List Categories and Criteria. Version 13. Prepared by the Standards and Petitions Subcommittee. Available ar http://cmsdocs.s3.amazonaws.com/RedListGuidelines.pdf. Accessed May/2020.
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), considering only the extent of occurrence (EOO) since the area of occupancy (AOO) is very sensitive to georeferencing bias associated with few or no georeferenced collections, and collecting effort (Särkinen et al. 2018SÄRKINEN T, POCZAI P, BARBOZA GE, VAN DER WEERDEN GM, BADEN M & KNAPP S. 2018. A revision of the Old World Black Nightshades (Morelloid clade of Solanum L., Solanaceae). PhytoKeys 106: 1-223., Knapp et al. 2019KNAPP S, BARBOZA GE, BOHS L & SÄRKINEN T. 2019. A revision of the Morelloid Clade of Solanum L. (Solanaceae) in North and Central America and the Caribbean. PhytoKeys 123: 1-144.). Geospatial Conservation Assessment Tool (GeoCAT) was used (Bachman et al. 2011BACHMAN S, MOAT J, HILL AW, DE LA TORRE J & SCOTT B. 2011. Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool. ZooKeys 150: 117-126.).
RESULTS
Solanaceae diversity in South America
The number of Solanaceae species in South America accounted is 1611 (1595 native and 16 introduced species; Table SI Table SI, Table SII, Table SIII. - Supplementary Material). The number of genera is 62 (61 native and one introduced, Datura L.) and the 10 most speciose genera are: Solanum, Cestrum L., Nolana L.f., Jaltomata Schltdl., Lycianthes (Dunal) Hassl., Deprea Raf., Nicotiana L., Iochroma Benth., Lycium, and Capsicum L. (Fig. 2; Table SII Table SI, Table SII, Table SIII. - Supplementary Material). Analysis of species richness across South American countries shows that Peru has the highest number of genera, species as well as endemic species (Table I, Table SIII Table SI, Table SII, Table SIII. -Supplementary Material) while Uruguay and the Guianas have the lowest numbers. Paraguay and the Guianas have only one endemic species (Cestrum hassleri Francey and Solanum costatum M.Nee, respectively). Chile, Peru and Brazil are the countries with the greatest percentage of endemic species (almost 50%).
Distribution of Solanaceae in South American countries. a. The 10 most speciose genera. b. The 10 most endemic speciose genera. Abbreviations: ARG (Argentina), BOL (Bolivia), BRA (Brazil), CHI (Chile), COL (Colombia), ECU (Ecuador), GUI (The Guianas), PAR (Paraguay), PER (Peru), URU (Uruguay), VEN (Venezuela).
Species diversity of Solanaceae in South America. Number of genera, species and endemics is provided for each country and the Guianas (French Guiana, Guyana and Suriname).
The relationship between the number of genera, species or endemics and the area of the countries was significantly positive (p≤0.05; R2 =0.36, R2 =0.44, R2 =0.45, respectively) (Fig. 3a-c). Ecuador and Peru show higher values than expected according to their areas in all cases and inversely, the Guianas and Paraguay (only for the case of endemics and area) show lower values than expected for their areas.
Solanaceae in South America: relationships between the number of taxa and the countries area, and between endemic species and total species. a. Number of genera and area. b. Number of species and area. c. Number of endemic species and area. d. Number of endemic species and total number of species for different countries. Abbreviations: Arg (Argentina), Bol (Bolivia), Bra (Brazil), Chi (Chile), Col (Colombia), Ecu (Ecuador), Gui (The Guianas), Par (Paraguay), Per (Peru), Uru (Uruguay), Ven (Venezuela).
Species richness was positively related to endemics (R2 =0.80, p <0.01), so the countries with a larger number of species also possess a larger number of endemic species (Fig. 3d). Chile and Brazil present a larger number of endemics than expected from the total number of species; inversely, Bolivia, Colombia, and Ecuador possess a smaller number.
Solanaceae diversity and distribution in Argentina
Argentine Solanaceae in numbers
Argentina has 338 specific and infraspecific taxa, distributed in 34 genera and 319 species of which 83 are endemics (Fig. 4a). Regarding their distribution status, almost all species are native to Argentina, except for seven species (Fig. 4b). Lycium barbarum L., from Asia, is the only extra-American species. The other introduced species are American: Datura ferox L., D. inoxia Mill., D. stramonium L. and Solanum rostratum Dunal from North America, and Brugmansia suaveolens (Willd.) Sweet and Solanum capsicoides All. from Brazil.
Solanaceae diversity in Argentina. a. Number of species, specific and infraspecific taxa, and endemics in each genus (taxa means: specific and infraspecific taxa). b. Number of specific and infraspecific taxa versus distribution status. c. Pantacantha, the only endemic monotypic genus in Argentina.
A total of 38% of the genera (13) is represented by only one species from which Pantacantha Speg. is the sole monotypic genus restricted to Argentina (Fig. 4c). Contrarily, the genus with the largest number of species is Solanum, also being the one that contributes with the most endemics in the country. Considering the genera with more than three species, those with the greatest proportion of endemics are Sclerophylax with 86% and Benthamiella Speg. with 67% (Fig. 4a).
Argentine Solanaceae in ecoregions
Solanum and Lycium are present in all Argentine ecoregions; Nicotiana, Nierembergia Ruiz & Pav., Cestrum and Physalis L. also inhabit nearly all ecoregions, except for the Wet Temperate Pacific; Exodeconus Raf., Reyesia Gay and Salpiglossis are exclusive to the Andes, Athenaea Sendtn. and Brugmansia to the Atlantic Forest, Eriolarynx (Hunz.) Hunz. to Yungas and Melananthus Walp. to the Chaco. Benthamiella is a Chilean-Argentine Patagonian endemic genus. The ecoregions with the greatest richness of genera are the Chaco (with 65% of the genera i.e., 22 genera) and Yungas (62% i.e., 21 genera), followed by The Andes (20 genera), Monte (19 genera), Atlantic Forest and Pampas (18 genera), whereas Patagonia (44% i.e., 15 genera) and Wet Temperate Pacific (6% i.e., 2 genera) have the lowest richness.
The Chaco is the ecoregion with the highest number of specific and infraspecific taxa (121), followed by the Andes (111), Pampas (109), Monte (94), Yungas (90), Atlantic Forest (74), Patagonia (59) and Wet Temperate Pacific (5) (Fig. 5a). Introduced species are found in all regions but Wet Temperate Pacific.
Solanaceae diversity in Argentine ecoregions. a. Total number of specific and infraspecific taxa in each ecoregion. Percentage relative to the total Argentine specific and infraspecific taxa are expressed. b. Number of Argentine endemic specific and infraspecific taxa in each ecoregion. Percentage relative to the total Argentine endemic specific and infraspecific taxa are expressed.
The Argentine endemic specific and infraspecific taxa are recorded in six ecoregions: Monte (37), Andes (34), Chaco (29), Patagonia (29), Yungas (16), and Pampas (5) (Fig. 5b).
The analysis of the similarities and differences of Solanaceae between ecoregions showed that about 60% of the total variation is related to the number of genera, where Atlantic Forest, Pampas, Chaco and Yungas constitute a group on the lower left side of the biplot (Fig. 6a) with Monte related to them. When considering the total specific and infraspecific taxa, 40% of the total variability between ecoregions could be explained, and in this way Atlantic Forest, Pampas, Chaco, and Yungas are separated from the remaining regions (Fig. 6b), and the Pacific region being isolated from the rest. Regarding endemics (Fig. 6c), Monte and the Andes are the most similar ecoregions, sharing a high number of specific and infraspecific endemic taxa.
PCoA dispersion diagrams of Solanaceae species richness of Argentina (see Methods for details). a. Total genera (34). b. Total specific and infraspecific taxa (338). c. Total endemic specific and infraspecific taxa (93).
Conservation status assessment of Argentine endemic species
Using the IUCN criteria, we consider that 34% (28) of the 83 endemic species belong to threatened categories (Fig. 7); being 9 Critically Endangered (CR), 6 Endangered (EN), and 13 Vulnerable (VU). Six species are considered as Near Threatened (NT) while 2 are Data Deficient (DD) (Fig. 8). The remaining species (47) were evaluated as Least Concern (LC).
Conservation status of Solanaceae species endemic to Argentina. The IUCN criteria used for classification are indicated. Asterisk (*) indicates that the best estimates of population size is 2,000 mature individuals, but this estimate is very uncertain, and as low as 1,000 mature individuals cannot be ruled out.
Data Deficient species from Argentina. a. Jaborosa chubutensis (taken from Barboza & Hunziker 1987BARBOZA GE & HUNZIKER AT. 1987. Estudios sobre Solanaceae XXV. Revisión de Jaborosa. Kurtziana 19(1): 77-153.). b. Lycium pubitubum (taken from Bernardello 1986BERNARDELLO G. 1986. Revisión taxonómica de las especies sudamericanas de Lycium (Solanaceae). Bol Acad Nac Ci Córdoba 57(3-4): 173-356.).
DISCUSSION
The spatial patterns in species richness of Solanaceae in the South American countries agree with the species-area relationships found by Cowling & Samways (1994)COWLING RM & SAMWAYS MJ. 1994. Predicting global patterns of endemic plant species richness. Biodivers Lett 2(5): 127-131. and Gaston (2000)GASTON KJ. 2000. Global patterns in biodiversity. Nature 405(6783): 220-227.. Brazil, the most extensive country, is also the most diverse in its vascular flora as well as the one with the greatest proportion of endemics (Ulloa Ulloa et al. 2017ULLOA ULLOA C ET AL. 2017. An integrated assessment of the vascular plant species of the Americas. Science 358(6370): 1614-1617.). Peru and Ecuador present higher Solanaceae diversity than expected. According to our results, Peru is the country with the greatest diversity including genera, total species and endemic species. Solanum, Jaltomata and Nolana are the most speciose genera in Peru (Knapp et al. 2006KNAPP S, SPOONER DM & LEÓN B. 2006. Solanaceae endémicas del Perú. Rev Peru Biol 13(2): 612-643.), and Peru specially holds the greater number of potato (Spooner et al. 2016SPOONER DM, ALVAREZ N, PERALTA IE & CLAUSEN AM. 2016. Taxonomy of wild potatoes and their relatives in Southern South America (Solanum sect. Petota and Etuberosum). Syst Bot Monogr 100: 1-240.) and tomato species in the world (Peralta et al. 2008PERALTA IE, SPOONER DM & KNAPP S. 2008. Taxonomy of wild tomatoes and their relatives (Solanum sect. Lycopersicoides, sect. Juglandifolia, sect. Lycopersicon; Solanaceae). Syst Bot Monogr 84: 1-186.). On the other hand, Solanum, Cestrum and Deprea are the most speciose genera in Ecuador (see Table SIII).
In addition, Chile and Brazil have a high proportion of Solanaceae endemic species; they even have endemic genera such as Vestia Willd. and Latua Phil. in Chile, and Duckeodendron, Dyssochroma Miers, Heteranthia Nees & Mart. and Metternichia J.C.Mikan in Brazil. Both countries have also a high percentage of endemic species considering all vascular plants: 55% for Brazil (Flora do Brasil 2019bFLORA DO BRASIL. 2019b. Flora do Brasil 2020 em construção. Jardim Botânico do Rio de Janeiro. Disponível em http://floradobrasil.jbrj.gov.br/. Acessado em Maio/2020.
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) and almost 40% for Chile (Rodriguez et al. 2018RODRIGUEZ R ET AL. 2018. Catálogo de las plantas vasculares de Chile. Gayana Bot 75(1): 1-430.).
The non South American countries, Costa Rica and Mexico, have the highest biodiversity indexes in the world (Grayum et al. 2004GRAYUM MH, HAMMEL BE, TROYO S & ZAMORA N. 2004. Historia: La exploración botánica y la florística en Costa Rica. In: Hammel BE, Grayum MH, Herrera C & Zamora N (Eds), Manual de plantas de Costa Rica, 1st ed., Monogr Syst Bot Missouri Bot Gard 97: 1-48., Villaseñor 2016VILLASEÑOR JL. 2016. Checklist of the native vascular plants of Mexico. Revista Mex Biodivers 87(3): 559-902.); specifically, Mexico has a great Solanaceae diversity, with 34 genera and 381 species (Martínez et al. 2017MARTÍNEZ M, VARGAS-PONCE O, RODRÍGUEZ A, CHIANG F & OCEGUEDA S. 2017. Solanaceae family in Mexico. Bot Sci 95(1): 131-145.), while in Costa Rica the number is lower (23 genera and 180 species) (Bohs 2015BOHS L. 2015. Solanaceae. In: Hammel BE, Zamora N & Grayum MH (Eds), Manual de Plantas de Costa Rica, Vol. VIII, 1st ed., Monogr Syst Bot Missouri Bot Gard 131: 205-336.). Thus, the amount of these taxa for each country match with what is expected according to their areas, as happens in several South American countries (e.g., Argentina, Brazil, Colombia, Venezuela) tested in this paper.
Regarding Argentina, our inventory contributed to the increase of number of species compared to Barboza (2013)BARBOZA GE (Coord). 2013. Solanaceae. In: Zuloaga FO, Belgrano M & Anton AM (Eds), Flora Argentina, 1st ed., San Isidro: IBODA-IMBIV, CONICET 13: 1-350. adding one species (i.e., 319), while the number of specific and infraspecific taxa reminded at 338. Solanum rostratum is the new introduced species (Del Vitto & Petenatti 2015DEL VITTO LA & PETENATTI EM. 2015. Sobre la presencia de Solanum rostratum (Solanaceae) en Sudamérica: una neófita tóxica de gran potencial como maleza. Rev Fac Cienc Agrar Univ Nac Cuyo 47(2): 109-121.). Beyond the numbers, it should be noted that there were some changes in the taxonomy and distribution: Solanum chamaesarachidium Bitter and Solanum excisirhombeum Bitter are now considered synonyms of Solanum weddellii Phil. and Solanum grandidentatum Phil (Solanaceae Source 2019SOLANACEAE SOURCE. 2019. A global taxonomic resource for the nightshade family. Available at http://www.solanaceaesource.org/. Accessed on May/2020.
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), respectively; Salpichroa ramosissima Miers was excluded from the Argentinean flora, Salpichroa tristis var. lehmanni (Dammer) Keel is now considered at specific level (i.e., Salpichroa lehmannii Dammer) (Gonzáles et al. 2018GONZÁLES P, BASSO AV, SÄRKINEN T, LEIVA GONZÁLEZ S, CANO A & BARBOZA GE. 2018. Diversidad y distribución del género Salpichroa (Solanaceae), con énfasis en los Andes peruanos. Darwiniana NS 6(1): 24-34.); and the genus Iochroma does not grow in Argentina anymore since Iochroma australe Griseb has been recently transferred to Eriolarynx (Eriolarynx australis (Griseb.) J.M.H. Shaw) (Shaw 2018SHAW J. 2018. Iochroma reshuffle. Plantsman 17(2): 124-125.).
The Yungas and Atlantic Forest are the ecoregions of greatest total species richness of vascular plants (Grossi et al. 2012GROSSI MA, GUTIÉRREZ DG & DELUCCHI G. 2012. Una mirada sobre el estado actual de la conservación de la flora argentina. Conserv Veg 16: 15-17., Zuloaga & Belgrano 2015ZULOAGA FO & BELGRANO MJ. 2015. The Catalogue of Vascular Plants of the Southern Cone and the Flora of Argentina: their contribution to the World Flora. Rodriguésia 66(4): 989-1024.). However, for Solanaceae, the Chaco is the ecoregion with the largest number of genera, and specific and infraspecific taxa. For endemics, Monte is the ecoregion with the largest number of endemic specific and infraspecific taxa of Solanaceae, coinciding with southern South America total endemic vascular plants (Zuloaga & Belgrano 2015ZULOAGA FO & BELGRANO MJ. 2015. The Catalogue of Vascular Plants of the Southern Cone and the Flora of Argentina: their contribution to the World Flora. Rodriguésia 66(4): 989-1024.). The Andes and Patagonia, important ecoregions for total endemic vascular species, particularly the Poaceae and Asteraceae (Katinas et al. 2007KATINAS L, GUTIÉRREZ DG, GROSSI MA & CRISCI JV. 2007. Panorama de la familia Asteraceae (= Compositae) en la República Argentina. Bol Soc Argent Bot 42(1-2): 113-129., Biganzoli & Zuloaga 2015BIGANZOLI F & ZULOAGA FO. 2015. Analysis of Poaceae biodiversity in austral South America. Rodriguésia 66(2): 337-351.), are also main centres for endemics Solanaceae (see Fig. 5b).
Although in this work environmental aspects were not included in the diversity analyses, some general remarks could be outlined. Some genera appear to have specific habitat or bioclimatic preferences in their distribution in South America. Many of the species of Calibrachoa Cerv. have a very limited geographical distribution and, although some have overlapping occurrence areas, they generally occupy different environments, characterized by different types of soils in general with acid pH (Greppi et al. 2013GREPPI JA, HAGIWARA JC & STEHMANN JR. 2013. Novelties in Calibrachoa (Solanaceae) and taxonomic notes on the genus for Argentina. Darwiniana, NS 1(1): 173-187.). Solanum diversity patterns have been widely explored showing higher species richness in mountains, and hence high diversity in Andean countries (e.g. Särkinen et al. 2015SÄRKINEN T ET AL. 2015. Listado anotado de Solanum L. (Solanaceae) en el Perú. Rev Peru Biol 22(1): 3-62.). Nolana species are largely restricted to fog-dependent desert lomas formations of coastal Peru and Chile (Dillon et al. 2009DILLON MO, TU T, XIE L, QUIPUSCOA SILVESTRE V & WEN J. 2009. Biogeographic diversification in Nolana (Solanaceae), a ubiquitous member of the Atacama and Peruvian Deserts along the western coast of South America. J Syst Evol 47(5): 457-476.). Four endemic Brazilian genera live in restricted areas: Dyssochroma inhabits exclusively in the Atlantic Forest, Heteranthia and Metternichia grow in the Caatinga and Atlantic Forest while Duckeodendron is from Amazonia (Flora do Brasil 2019aFLORA DO BRASIL. 2019a. Solanaceae in Flora do Brasil 2020 em construção. Jardim Botânico do Rio de Janeiro. Disponível em http://floradobrasil.jbrj.gov.br/reflora/floradobrasil/FB225. Acessado em Maio/2020.
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). Finally, genera of the tribe Benthamielleae (Hunz.) Hunz. are typical of Patagonia, where Pantacantha is mostly an orophilous genus (Hunziker 2001HUNZIKER AT. 2001. The genera of Solanaceae, 1st ed., Ruggell: A.R.G. Gantner Verlag K.-G, p. 1-500., Barboza GE & Cantero JJ, personal communication).
Among the native species growing in Argentina, some of them have small or infrequent populations or there are no recent collections; these are: Calibrachoa pubescens (Spreng.) Stehmann, Cestrum bracteatum Link & Otto, C. reflexum Sendtn., Exodeconus integrifolius (Phil.) Axelius, Lycium cyathiforme C.L. Hitchc., Melananthus multiflorus Carvalho, Salpiglossis sinuata Ruiz & Pav., Schizanthus hookeri Gillies ex Graham, Sessea vestioides (Schltdl.) Hunz., S. regnellii Taub., Solanum furcatum Dunal, S. grandidentatum Phil., S. toldense Matesevach & Barboza and S. turneroides Chodat (Barboza 2013BARBOZA GE (Coord). 2013. Solanaceae. In: Zuloaga FO, Belgrano M & Anton AM (Eds), Flora Argentina, 1st ed., San Isidro: IBODA-IMBIV, CONICET 13: 1-350.). Mostly these species seem to be locally rare or really discrete, and therefore hard to find. One example is the endemic Jaborosa ameghinoi (Speg.) Macloskie & Dusén, rediscovered by González et al. (2017)GONZÁLEZ C, LLORENS M, CALDERÓN A, SIMÓN P, SILVA C, PONCE G & PERAL M. 2017. Rediscovery and characterización of Jaborosa ameghinoi, through over a century, in the Austral Mount of Patagonia, Argentina. Bol Soc Argent Bot 52(Suppl.): 277. after more than a century since the original collection, which was probably overlooked many times because it loses its aerial parts and stays under the soil during most of the year, as happens in other Patagonian species (e.g., as other Jaborosa species or Fabiana nana (Speg.) S.C. Arroyo; Barboza GE, personal communication). We recommend carrying out further sampling efforts that could improve the knowledge of species’ geographic distribution which is essential for conservation planning and forecasting (Elith et al. 2006ELITH J ET AL. 2006. Novel methods improve prediction of species’ distributions from occurrence data. Ecography 29(2): 129-151.). In relation to the introduced Solanaceae species in Argentina, the Asiatic Lycium barbarum is cultivated in Santa Cruz province where it could escape from gardens (Bernardello 2013BERNARDELLO G. 2013. Lycium L. In: Zuloaga FO, Belgrano M & Anton AM (Eds), Flora Argentina, Solanaceae, 1st ed., San Isidro: IBODA-IMBIV, CONICET, 13: 47-75.). Usually, the North American species of Datura (D. ferox, D. inoxia and D. stramonium) grow as weeds and ruderals, mainly D. ferox which is an important noxious plant in summer crop fields (Fernández et al. 2016FERNÁNDEZ O, LEGUIZAMÓN ES, ACCIARESI HA, TROIANI HO & VILLAMIL CB. 2016. Malezas e Invasoras de la Argentina. Tomo II: Descripción y Reconocimiento, 1st ed., Bahía Blanca: Ediuns, p. 1-936.). Up to now, the Brazilian Brugmansia suaveolens (Hay 2014HAY A. 2014. Brugmansia suaveolens. The IUCN Red List of Threatened Species 2014: e.T51247699A58913403. Available at https://www.iucnredlist.org/es/species/51247699/58913403/. Accessed on May/2020.
https://www.iucnredlist.org/es/species/5...
, Dupin 2017DUPIN J. 2017. Historical Biogeography and the Evolution of Environmental Niche and Fruit Type in Datureae (Solanaceae). Ecology & Evolutionary Biology Graduate Theses & Dissertations. Available at https://scholar.colorado.edu/ebio_gradetds/102 .Accessed on May/2020.
https://scholar.colorado.edu/ebio_gradet...
) has been collected infrequently at different sites in NE Argentina (Misiones) (Documenta Florae Australis 2019DOCUMENTA FLORAE AUSTRALIS. 2019. Available at http://www.darwin.edu.ar/iris/ .Accessed on May/2020.
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) and Solanum capsicoides is cultivated as ornamental (Chiarini 2013CHIARINI FE. 2013. Grupo VI. Leptostemonum. In: Zuloaga FO, Belgrano M & Anton AM (Eds), Flora Argentina: Vol 13, Solanaceae, 1st ed., San Isidro: IBODA-IMBIV, CONICET, p. 206-231.). One population of the North American Solanum rostratum was recently found in central western Argentina (San Luis Province) as a ruderal (Del Vitto & Petenatti 2015DEL VITTO LA & PETENATTI EM. 2015. Sobre la presencia de Solanum rostratum (Solanaceae) en Sudamérica: una neófita tóxica de gran potencial como maleza. Rev Fac Cienc Agrar Univ Nac Cuyo 47(2): 109-121.) and no other record has been cited.
Only a few species of Solanaceae were previously assigned to IUCN categories in Argentina; Physalis victoriana J.M. Toledo was considered as Endangered (EN) by Toledo (2013)TOLEDO JM. 2013. Physalis victoriana (Solanaceae) a new species from Northern Argentina. Phytotaxa 124(1): 60-64. and Solanum pygmaeum Cav. as Least Concern (LC) by Särkinen et al. (2018)SÄRKINEN T, POCZAI P, BARBOZA GE, VAN DER WEERDEN GM, BADEN M & KNAPP S. 2018. A revision of the Old World Black Nightshades (Morelloid clade of Solanum L., Solanaceae). PhytoKeys 106: 1-223., in coincidence with our proposal. In another case, Delucchi (2006)DELUCCHI G. 2006. Las especies vegetales amenazadas de la Provincia de Buenos Aires: Una actualización. APRONA Bol Cient 39(1): 19-31. suggests a local categorization for plants of Buenos Aires, categorizing the endemic Nierembergia tandilensis (Kuntze) Cabrera as Critically Endangered (CR); however, we propose to reformulate its status as Vulnerable (VU) at global scale. A constant update of the conservation status of the species is necessary, since population changes can happen (IUCN 2017). Furthermore, efforts to recollect the two endemics with Data Deficient (DD), Jaborosa chubutensis Barboza & Hunz. and Lycium pubitubum C.L. Hitchc., only known by their type collections (Bernardello 1986BERNARDELLO G. 1986. Revisión taxonómica de las especies sudamericanas de Lycium (Solanaceae). Bol Acad Nac Ci Córdoba 57(3-4): 173-356., Barboza & Hunziker 1987BARBOZA GE & HUNZIKER AT. 1987. Estudios sobre Solanaceae XXV. Revisión de Jaborosa. Kurtziana 19(1): 77-153.), are needed.
Land use change, mainly deforestation, causes environmental impacts such as biodiversity loss in South America (De Sy et al. 2015DE SY V, HEROLD M, ACHARD F, BEUCHLE R, CLEVERS JGPW, LINDQUIST E & VERCHOT L. 2015. Land use patterns and related carbon losses following deforestation in South America. Environ Res Lett 10(12): 124004.). The expansion of the agricultural frontier has promoted deforestation in tropics. Brazil, Argentina and Paraguay are the three countries with the largest loss of tree cover during 1982-2016, and Argentina alone lost an area of 113,000 km2 (25%) during these years (Song et al. 2018SONG XP, HANSEN MC, STEHMAN SV, POTAPOV PV, TYUKAVINA A, VERMOTE EF & TOWNSHEND JR. 2018. Global land change from 1982 to 2016. Nature 560(7720): 639-643.). In Argentina, during the 20th century and especially in more recent decades, the lowland landscape occupied by Chaco and Espinal forests has been transformed into an agricultural system at high deforestation rates (Boletta et al. 2006BOLETTA PE, RAVELO AC, PLANCHUELO AM & GRILLI M. 2006. Assessing deforestation in the Argentine Chaco. Forest Ecol Manag 228(1-3): 108-114.). In the mountain area of the same regions, fire, grazing and the recent advance of exotic woody species have transformed the landscape into a complex mosaic of degraded native forests, monospecific stands of woody exotics, shrublands and grasslands (Cabido et al. 2018CABIDO M, ZEBALLOS SR, ZAK M, CARRANZA ML, GIORGIS MA, CANTERO JJ & ACOSTA ATR. 2018. Native woody vegetation in central Argentina: Classification of Chaco and Espinal forests. Appl Veg Sci 21(2): 298-311.). In relation to these changes in land use, it is alarming that many of the taxa proposed here as threatened are only found in the margins of cultivated areas, increasing the risk of loss of their populations as a consequence of habitat changes (i.e., Solanum concarense Hunz.). Godoy-Bürki et al. (2014)GODOY-BÜRKI AC, ORTEGA-BAES P, SAJAMA JM & AAGESEN L. 2014. Conservation priorities in the Southern Central Andes: mismatch between endemism and diversity hotspots in the regional flora. Biodivers Conserv 23(1): 81-107. showed that in north-western (NW) Argentina, most protected areas are concentrated in the ecoregion with the highest diversity, the humid Yungas region, but the endemics are mainly found in the arid NW Argentina ecoregions. They also considered that the current reserve system in NW Argentina is ineffective. In our case, most of Solanaceae endemics grow in the Monte and Andes arid ecoregions also pointing out the need for choosing new priority areas for conservation as it has already been stated by Barboza et al. (2016)BARBOZA GE, CANTERO JJ, CHIARINI FE, CHIAPELLA J, FREIRE S, NUÑEZ CO, PALCHETTI V & ARIZA ESPINAR L. 2016. Vascular plants of Sierra de Famatina (La Rioja, Argentina): an analysis of its biodiversity. Phytotaxa 248(1): 1-123.. No formal program has been yet established for conservation of the endemic flora in Argentina. Only a preliminary data base (PlanEAr 2019PLANEAR - Plantas Endémicas de la Argentina. 2019. Available at http://www.lista-planear.org/. Accessed on May/2020.
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) for endemic species was generated recently, which is not updated nor based on IUCN criteria.
In Argentina, scientific research has been historically supported by public funds and the national state usually defines research priority areas and programs (Bekerman 2016BEKERMAN F. 2016. El desarrollo de la investigación científica en Argentina desde 1950: entre las universidades nacionales y el Consejo Nacional de Investigaciones Científicas y Técnicas. Rev Iberoam Educ Super 7(18): 3-23.). In this sense, the Red List of vascular plants in Argentina should be a priority issue in the near future.
CONCLUSIONS
South America has a great diversity of Solanaceae which represents approximately 60% of the Solanaceae diversity at the global scale and it is extremely variable between South American countries. Peru and Brazil host the highest diversity and the highest number of endemic species of Solanaceae, so its conservation should be a priority issue considering its biological and economic importance.
In Argentina, Chaco is the most diverse ecoregion for Solanaceae while Monte and Andes have the highest number of endemic species. Furthermore, the conservation status assessment revealed that 34% of the endemic species of Solanaceae belongs to threatened categories. No formal conservation program exists for Argentine vascular plants. We propose the Monte and Andes ecoregions as priority areas for conservation and an urgent agenda for the Chaco ecoregion where land use changes have dramatically destroyed the natural habitats where a high percentage of endemic Solanaceae grow.
ACKNOWLEGMENTS
The authors thanks to curators and staff of the Argentine herbaria for their disposition to provide information of the Solanaceae collections; to G. Bernardello, F. Chiarini, A.A. Cocucci, R. Deanna, L. Giacomin, M. Matesevach, and A. Romanutti for specifying either the identification or the distribution and the abundance of the populations of some species; to R. Deanna for Nierembergia rigida and Physalis viscosa photographs; and to the anonymous reviewers for constructive suggestions to improve the manuscript. V.P. thanks the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET) for the doctoral fellowship. This study was funded by the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET)-Argentina (PIP number 11220170100147CO), Secretaría de Ciencia y Tecnología (SECyT)-UNC (Res. 411-18, Universidad Nacional de Córdoba, Argentina) and SECyT-UNRC (Res. Rec. 130/2017, Universidad Nacional de Río Cuarto, Argentina).
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SUPPLEMENTARY MATERIAL
Table SI, Table SII, Table SIII.
Publication Dates
-
Publication in this collection
07 Aug 2020 -
Date of issue
2020
History
-
Received
7 Jan 2019 -
Accepted
14 Oct 2019